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To investigate the phylogeographical patterns of red deer (Cervus elaphus) in Europe, and to disentangle the influence of ancient (e.g. Pleistocene ice ages) from more recent processes (e.g. human translocations). Europe. In this study we provide by far the most extensive analysis of genetic structure in European red deer, based on analyses of variation at two mitochondrial markers (cyt b and D-loop) in a large number of individuals from 39 locations. Relationships of mitochondrial DNA haplotypes were determined using minimum spanning networks and phylogenetic analyses. Population structure was examined by analyses of molecular variance. Historical processes shaping the present patterns were inferred from nested clade analysis and nucleotide diversity statistics. Within Europe, we detected three deeply divergent mitochondrial DNA lineages. The three lineages displayed a phylogeographical pattern dividing individuals into western European, eastern European and Mediterranean (Sardinia, Spain and Africa) groups, suggesting contraction into three separate refugia during the last glaciation. Few haplotypes were shared among these three groups, a finding also confirmed by FST values. Calculations of divergence times suggest that the groups probably split during the Pleistocene. The observed pattern is interpreted to result from isolation in different refugia during the last glaciation. The western and eastern European lineages could be linked to an Iberian and Balkan refugium, respectively. The third lineage might originate from a Sardinian or African refugium. We link local phylogeographical patterns observed in Europe to the post-glacial recolonization process, shaped by the geographical localization of refugia and barriers to gene flow. Regardless of the importance of red deer as a game species and the tradition of translocating red deer in Europe, we detected few individuals that did not match the trichotomous pattern, suggesting that translocations have occurred mainly at smaller spatial scales.

Even though theoretical models show that parasites may regulate host population densities, few empirical studies have given support to this hypothesis. We present experimental and observational evidence for a host-parasite interaction where the parasite has sufficient impact on host population dynamics for regulation to occur. During a six year study of the Svalbard reindeer and its parasitic gastrointestinal nematode Ostertagia gruehneri we found that anthelminthic treatment in April-May increased the probability of a reindeer having a calf in the next year, compared with untreated controls. However, treatment did not influence the over-winter survival of the reindeer. The annual variation in the degree to which parasites depressed fecundity was positively related to the abundance of O. gruehneri infection the previous October, which in turn was related to host density two years earlier. In addition to the treatment effect, there was a strong negative effect of winter precipitation on the probability of female reindeer having a calf. A simple matrix model was parameterized using estimates from our experimental and observational data. This model shows that the parasite-mediated effect on fecundity was sufficient to regulate reindeer densities around observed host densities.

1. It is well known that gastrointestinal nematodes can have a significant impact on the growth of farmed ruminants. The clinical signs of these parasites are often subtle, with production losses mainly due to reductions in host appetite and gut function. However, little is known about the impact of this widespread group of parasites on wild ruminants. We use experiments and cross-sectional data to investigate the effects of gastrointestinal nematodes on a wild host, the Svalbard reindeer. 2. Individually marked reindeer were treated for parasites. Their body mass, back fat depth and fecundity were compared with the control group one year later. The effect of treatment on worm burdens was investigated in a subsample of animals that were culled 3 and 6 months after treatment. Also, the relationship between the intensity of infection and body mass, back fat depth and host pregnancy rates was investigated in cross-sectional data from culled reindeer. 3. The anthelmintic treatment caused an increase in the body mass, back fat depth and fecundity of the reindeer. Treatment depressed the abundance of adult parasites of Ostertagia gruehneri for at least 6 months, but had no significant effect on the abundance of adults of the other dominant parasite species, Marshallagi marshalli, or the abundance of larval stages of either species. 4. In the cross-sectional study, the probability of a reindeer being pregnant in late winter was negatively related to the abundance of adult O. gruehneri when controlling for host body mass. However, no clear evidence were found for an effect of parasitism on host condition in the cross-sectional study. 5. Our experimental results show for the first time in a natural ruminant host population that gastrointestinal nematodes can have a significant effect on host condition and fecundity. However, effects of infection on host condition was not detectable in the cross-sectional study. Also, we show that the experimental effects on host condition and fecundity is most likely to be due to a negative effect of O. gruehneri, while the experimental design did not allow detection of potential negative effects of M. marshalli.

1. A central concept in ruminant foraging ecology is that even slight changes in plant quality affect body growth substantially, because ruminants not only gain more protein and energy but also use less time for rumination when eating high-quality forage. Increased access to highly nutritious forage is thus regarded as an important driving force in the evolution of migration in large herbivores, because the temporal and spatial variation in plant quality is huge. Body weight is in turn a major determinant of reproductive performance and survival in ungulates, and therefore important for population dynamics. 2. In the temperate region, the vertical movement of cervids from a low-elevation winter range to a high-elevation summer range is the most common pattern of migration. Migration to high elevations during summer is regarded as a strategy to increase energy intake among northern temperate ungulates. 3. It has been suggested that there is increased forage quality at high elevation. This leads to the prediction that body weights are positively correlated with the proportion of high-altitude habitat. Alternatively, cervids may benefit due to prolonged access to newly emerged forage as they migrate along an altitudinal gradient. If so, it can be predicted that body weights are highest in regions with the highest diversity of different altitudes. 4. The facing of slopes, i.e. the aspect, is also regarded as an important part of the habitat for deer. Access to north-facing slopes is regarded as favourable because plant quality may be higher on north-facing slopes; access to a variety of different aspects is also favourable because this may lead to a variable snow melt and thus a longer time period with access to high-quality forage. 5. In order to test the above predictions (under points 3 and 4), we analysed the spatial variability of body weight of 8452 female and 12474 male red deer harvested during 1975-98 from 105 municipalities in Norway. 6. Body weight of red deer was positively related to the diversity of altitudes, and negatively related to the proportion of high-altitude habitat per se. This demonstrated that the altitudinal migration of cervids is not necessarily due to increased forage quality at high elevation, but rather that migrating cervids gain from prolonged access to newly emerging forage along an altitudinal gradient as they migrate to high elevation during early summer. 7. There was also no support for the hypothesis that access to a high proportion of north-facing slopes was favourable; rather, body weight of red deer was correlated with access to a diversity of aspects. There was thus clear evidence that a variable topography, measured as different altitude levels and aspects, was positively related to body weight of red deer. 8. We discuss the possible causes of why an ideal free distribution is not reached and conclude that phenological differences in plants related to topography can have a large impact on body weights of cervids, and therefore induce considerable spatial heterogeneity in population dynamics.

There are only a few recent studies that have demonstrated senescence in ungulates and nothing is known regarding how patterns of senescence may vary as a function of density. Senescence in general is linked to the cost of reproduction, which probably increases with density in ungulates and may differ between the sexes. Further, senescence in ungulates is also regarded to be a function of tooth wear rates. As density dependence and sexual differences in food choice have been well documented, this may lead to different tooth wear rates and, thus, possibly density-dependent and sex-specific patterns of senescence. We therefore investigated the effects of age, sex, density and their possible interactions on the variability of body weight in 29 047 red deer harvested during 1965-1998 from Norway, out of which 380 males and 1452 females were eight years or older.There was clear evidence that spatio-temporal variation in density correlated negatively with body weights. In addition, there was evidence of senescence in both male and female red deer. Age at onset of senescence in females was after 20 years of age and independent of population density. In males, the onset and rate of senescence increased with increasing population density. The onset of senescence for males was at ca. 12 years of age at low density, but decreased to approximately ten years of age at high density. The pattern of density-dependent senescence in males, but not that in females, can be explained if (i) the cost of reproduction increases with density more strongly in male than in female red deer, and/or (ii) tooth wear rates are density dependent in males, but not in females. We discuss the ability of these two different, not mutually exclusive hypotheses in explaining the observed pattern of senescence.

1. Timing and synchrony of reproduction are regarded as crucially important factors for fitness in seasonal environments. Natural selection has probably favoured temperate and arctic female herbivores that match reproduction with onset of plant growth in spring. However, breeding synchrony may also be affected by variation in phenotypic quality of females in a population, because females in poor body condition have been found to delay ovulation and subsequent calving. 2. We compared breeding phenology, i.e. the timing and synchrony of rutting (roaring, sexual aggregation) and calving of red deer (Cervus elaphus L.) in France (latitude: 49°N) and Norway (latitude: 63°N). We hypothesized (H1) that calving and rutting were later at the site with latest onset of plant growth. 3. We further quantified overall environmental predictability as the sum of annual constancy and seasonality and tested three different (not mutually exclusive) hypotheses about breeding synchrony: (H2a) the population experiencing most seasonal plant phenology should show the highest breeding synchrony; (H2b) overall predictability of plant phenology should determine breeding synchrony; and (H2c) breeding should be more synchronized in the population with lowest female body weight variation within age classes because they ovulate more synchronously. 4. Calving and rutting, as well as onset of plant phenology, were later in Norway than in France, complying with the first hypothesis. Plant growth in spring was overall more predictable and also more seasonal in Norway than France. Hence we expected higher breeding synchrony in Norway than in France according to H2aand H2b. Variance in female body weight was slightly higher in France than in Norway, which should also cause more synchronized breeding in Norway than in France (H2c). Contrary to all predictions, variance in rutting and calving dates was around two times higher in Norway than in France. 5. We suggest two alternative explanations of breeding synchrony. A more variable topography in Norway can make optimal birth date more variable on a local scale than in France, thereby maintaining a higher genetic variance for calving date in Norwegian red deer. Further, population age structure may play a role, as ovulation varies according to female age. Clearly, processes of breeding synchrony are far more complex than previously realized.

Sexual segregation outside the mating season occurs in most species of sexually dimorphic ungulates and has been extensively described in the literature, but the mechanisms causing segregation are still debated. The detailed pattern of sexual segregation throughout the year has rarely been presented for mammals, and no study, to our knowledge, has used latitudinal-related variation in breeding phenology to shed light on the underlying mechanisms. Recent methodological developments have made it possible to quantify separate components of segregation (social, habitat) and activity synchrony in animal groups, but these major improvements have so far been little used. We observed European red deer year round at two widely different latitudes (France and Norway) and tested three different mechanistic hypotheses of segregation related to: (i) predation risk; (ii) body-size-related forage selection; and (iii) activity budget. Habitat segregation peaked during calving in both populations and dropped rapidly after calving. Females with calves were more segregated from males than were females without calves, pointing to a key role of anti-predator behaviour even though large predators are absent in France and extremely rare in Norway. However, at both sites individuals also grouped with their own sex within habitat types (i.e. social segregation), and individuals in mixed-sex groups were less synchronized in activity type than individuals in either unisex male or unisex female groups, suggesting that differences in activity budgets are involved. Social segregation peaked during calving and was lowest during the rut (indicating aggregation) in both populations; these activities occurred one month later in the Northern populations, corresponding well with known differences in breeding phenology. We conclude that latitude-dependent breeding phenology shapes the seasonal pattern of sexual segregation and that sexual segregation in ungulates has multiple causes.

Plant phenology of Luzula heathland plots in Spitsbergen (78°N) was manipulated by adding or removing snow, which altered the time for plots (2 m×2 m; n=10) to become snow-free. A 2-week difference in snowmelt, equivalent to approximately one-sixth of the growing season, was achieved between advanced (first to be snow-free) and delayed (last to be snow-free) treatments, which influenced plant biomass and plant quality. Nitrogen content of the forage species decreased with time after snowmelt, whereas C:N ratio increased. Manipulation of snowmelt led to a shift in \"phenological time\", without altering these plant quality parameters as such. Early in the growing season, Svalbard reindeer (Rangifer tarandus platyrhynchus) selected the advanced plots which had been snow-free for longest, presumably because of the greater biomass of both Luzula confusa and Salix polaris, major components of reindeer diet at that time of the year. Moreover, the proportion of live Luzula leaves was highest in advanced plots, relative to both unmanipulated control and delayed plots. In contrast, plant quality, measured as nitrogen content and C:N ratio of leaves, was lowest in the preferred plots. Phenolic content did not differ among treatments, and is therefore unlikely to play a role in reindeer selection for plots with early snowmelt. Unlike in temperate regions, where selection for plant quality seems to be of major importance, selection for plant quantity might be an outcome of generally low levels of plant biomass and high forage quality during the growing season in the high Arctic. Reindeer selection for high plant biomass is likely to lead to a more favourable nitrogen and energy return than selection for high plant quality.

Seasonal changes in crude protein content of graminoids and herbs grazed by red deer were monitored from 1 May, or as soon as snow melt exposed the vegetation, until 15 October at five sites along an altitudinal gradient from coast to inland. Crude protein declined exponentially with time at all sites, but declined most rapidly from initially higher values at inland locations at high elevations. As a result crude protein was positively correlated with altitude and distance from the coast in early summer and negatively correlated in autumn. The relationships between protein content, date and altitude were used to estimate the quality of the diet of twelve radio-collared female red deer that migrated to summer ranges in the mountains. Individual differences in body weight were significantly related to the estimated, mean crude protein in vegetation available during the summer. Constraints on the timing of migration to exploit the maximal protein concentrations at higher altitudes and the fitness benefits of adaptive ranging behaviour are discussed.

1. Sexual segregation occurs in most species of sexually dimorphic ungulates. At least five not mutually exclusive hypotheses have been formulated to explain patterns of social, habitat and spatial segregation; the indirect competition hypothesis (H1), the nutritional needs hypothesis (H2), the activity budget hypothesis (H3), the weather sensitivity hypothesis (H4), and the predation risk hypothesis (H5). 2. Each hypothesis has a unique set of predictions with respect to the occurrence of segregation, and how seasonality, density dependence and reproductive status affect sexual segregation. 3. We tested this set of predictions in order to separate the hypotheses H1-H5 for patterns of sexual segregation of the Svalbard reindeer based on 9 years data on seasonal estimates of spatial, habitat and social (i.e. grouping with their own sex) segregation in combination with resource selection functions. 4. Our results do not support that one single mechanism causes segregation. The activity budget hypothesis, the nutritional needs hypothesis and the weather sensitivity hypothesis were all partially supported, while the predation risk hypothesis was discarded for Svalbard reindeer because predators have been absent for at least 5000 years. Several mechanisms are thus interacting to explain the full-year pattern of sexual segregation and the combination of mechanisms varies among species and populations.